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The Development and Improvement of Instructions JOURNEYS WITHIN THE LEUCOPHOROPTERINI: REVISION OF THE TRIBE, GENERA AND SPECIES, AND DESCRIPTION OF NEW GENERA AND SPECIES FROM AUSTRALIA AND THE INDO-PACIFIC A Dissertation by KATRINA LOUISE MENARD Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY May 2011 Major Subject: Entomology Journeys within the Leucophoropterini: Revision of the Tribe, Genera and Species, and Description of New Genera and Species from Australia and the Indo-Pacific Copyright 2011 Katrina Louise Menard JOURNEYS WITHIN THE LEUCOPHOROPTERINI: REVISION OF THE TRIBE, GENERA AND SPECIES, AND DESCRIPTION OF NEW GENERA AND SPECIES FROM AUSTRALIA AND THE INDO-PACIFIC A Dissertation by KATRINA LOUISE MENARD Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Approved by: Chair of Committee, James Woolley Committee Members, Spencer Johnston Randall Schuh John Oswald James Manhart Head of Department, David Ragsdale May 2011 Major Subject: Entomology iii ABSTRACT Journeys within the Leucophoropterini: Revision of the Tribe, Genera and Species, and Description of New Genera and Species from Australia and the Indo-Pacific. (May 2011) Katrina Louise Menard, B.S., The College of William and Mary; M.S., Texas A&M University Chair of Advisory Committee: Dr. James Woolley The tribe Leucophoropterini (Miridae: Phylinae) is a diverse assemblage of primarily Indo-Pacific and Australian bugs which are united by simple, small genitalia and a trend towards ant-mimetic body forms. Previous to this work, the relationship of the Leucophoropterini to the other tribes of Phylinae, as well as the generic relationships within the lineage, was unresolved. Further, the characters initially proposed to unite the tribe are brought into question with the addition of several recently discovered taxa from Australia. The Leucophoropterini is first re-evaluated within a phylogenetic analysis of the subfamily Phylinae, using a combined molecular and morphological dataset to test the monophyly of the lineage, re-test the character synapomorphies supporting it, and to determine the closest relatives to the tribe. The molecular dataset includes 4 genes (COII, 16S, 28S, and 18S), and 123 morphological characters for 104 taxa, which is analyzed in a parsimony analysis using Tree analysis using New Technology [TNT], a iv model-based analysis in RAxML, and a Bayesian analysis in Mr. Bayes. All three methods resulted in phylogenetic trees with nearly identical generic and tribal groupings, and a lineage containing Pseudophylus Yasunaga, Decomia Poppius and Tuxedo Schuh being sister-group to the Leucophoropterini. With the closest relatives to the Leucophoropterini determined for outgroup selection, a generic revision of the tribe including both Australian and Indo-Pacific taxa is accomplished using 137 morphological characters and is analyzed in an un-weighted and implied weighted parsimony analysis using TNT for 86 leucophoropterine taxa. The Indo-Pacific taxa of Leucophoropterini are found to be related to the Australian Leucophoropterini, and at least two genera within the tribe (Sejanus Distant, Leucophoroptera Poppius) were found to be paraphyletic. Lastly, taxa are revised within the context of the generic-level phylogenetic analysis, with new genera and species from Australia and the Indo-Pacific being described. v DEDICATION I would like to dedicate this work to my parents, Jim and Joan Menard, for their never-ending support; my advisers Dr. Randall Schuh and Dr. Woolley for their continual support in helping me accomplish this goal, and Miguel Zarate for emotional support for the duration of the writing of this work. vi ACKNOWLEDGEMENTS I would like to thank my committee chair, Dr. Woolley, and my committee members, Dr. Schuh, Dr. Oswald, Dr. Manhart, and Dr. Johnston for their guidance and support throughout the course of this research. I also would like to thank Dr. Schaffner, Dr. Schwartz, Dr. Henry, and Dr. Cassis, for their assistance in mastering mirid taxonomy and morphology for the completion of this work. Further, I would also like to thank fellow Plant Bug Planetary Biodiversity Inventory members Dr. Tatarnic, Dr. Forero, Dr. Wyniger, Dr. Weirauch, Dr. Yasunaga and Dr. Konstantinov for specimens, support, and advisement. Also critical to my research was the Section 9 lab at the American Museum of Natural History, in particular Dr. Wheeler and Ellen Trimarco for allowing me to accomplish my molecular lab work in their lab facility. Thanks also go to my friends and colleagues, and the department faculty and staff, for making my time at Texas A&M University a great experience. In particular, I would like to thank Ed Riley, Christine Johnson, and Geoff Thompson for helping with the specimen collecting and transfers between the various institutions during my research. I also want to extend my gratitude to the Woolley Lab for their support and encouragement during the last months of my research. Finally, thanks to my mother and father for their encouragement and support without which this would not have been possible. vii TABLE OF CONTENTS Page ABSTRACT.............................................................................................................. iii DEDICATION .......................................................................................................... v ACKNOWLEDGEMENTS ...................................................................................... vi TABLE OF CONTENTS.......................................................................................... vii CHAPTER I INTRODUCTION…............................................................................ 1 II TRIBAL LIMITS WITHIN THE SUBFAMILY PHYLINAE (HETEROPTERA: MIRIDAE), WITH A FOCUS ON THE TRIBE LEUCOPHOROPTERINI…................................................................ 2 Introduction.................................................................................... 2 Methods.......................................................................................... 6 Morphological Character Discussion............................................. 8 Phylogenetic Methods.................................................................... 45 Results............................................................................................ 47 Discussion...................................................................................... 49 Conclusion...................................................................................... 69 III A PHYLOGENETIC STUDY OF THE GENERIC RELATIONSHIPS WITHIN LEUCOPHOROPTERINI SCHUH (MIRIDAE: PHYLINAE).................................................................... 74 Introduction.................................................................................... 74 Methods.......................................................................................... 78 Results............................................................................................ 80 Discussion...................................................................................... 85 Conclusion...................................................................................... 116 IV THE LEUCOPHOROPTERINI OF AUSTRALIA AND THE INDO- PACIFIC: REVISION AND KEY TO THE CURRENTLY RECOGNIZED GENERA OF LEUCOPHOROPTERINI, REDESCRIPTION OF THE AUSTRALIAN FAUNA, AND viii CHAPTER Page DESCRIPTIONS OF NEW GENERA AND SPECIES………………………………………………………. ......... 119 Introduction .................................................................................... 119 Materials and Methods................................................................... 119 Key to Genera of Leucophoropterini ............................................. 121 Diagnoses and Descriptions of Genera and Species ...................... 126 Taxa Removed from Leucophoropterini ........................................ 422 Species Moved into Other Genera of Phylini................................. 426 Species Incertae Sedis .................................................................... 427 V SUMMATION………………………………………………………. 438 REFERENCES.......................................................................................................... 441 APPENDIX 1 ............................................................................................................ 450 APPENDIX 2 ............................................................................................................ 451 APPENDIX 3 ............................................................................................................ 493 APPENDIX 4 ............................................................................................................ 589 VITA ......................................................................................................................... 622 1 CHAPTER I INTRODUCTION The family Miridae (Hemiptera: Heteroptera), or plant bugs, is one of the largest families of true bugs with over 10,000 described species (Schuh 2008) and potentially thousands more undescribed. Two of the largest subfamilies within this family are the sister-groups Orthotylinae and Phylinae. The focus of this study is on the subfamily Phylinae, and in particular the tribe Leucophoropterini. The Leucophoropterini is a primarily Indo-Pacific and Australian lineage with some genera that are cosmopolitan, Old-World tropical,
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